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THIS  BOOK  IS  DUE  ON  THE  DATI 
INDICATED  BELOW  AND  IS  SUB 
JECT  TO  AN  OVERDUE  FINE  Ai 
POSTED  AT  THE  CIRCULATIO> 
DESK. 


D.  H.  HILL  LIBRARY 
N.  C.  STATE  UNIVERSITY 


50M/2-78 


NC: 


LiOrary 


naicJign 


Inheritance  in  Canaries 


BY 


CHARLES   B.   DAVENPORT, 

Director  of  the  Station  for  Experimental  Evolution, 
Carnegie  Institution  of  Washington. 


WASHINGTON.  D.  C. 

Published  by  the  Carnegie  Institution  of  Washington. 

1908. 


North  Carolina  State  Library 


Gift  of 


«i2^ 


North  Caress  State  Library 

Inheritance  in  Canaries 


BY 


CHARLES   B.   DAVENPORT, 

Director  of  the  Station  for  Experimental  Evolution, 
Carnegie  Institution  of  Washington. 


WASHINGTON,  D.  C. 
Published  by  the  Carnegie  Institution  of  Washington. 

1908. 


Carnegie  Institution  of  Washington  Publication  No.  95. 
Papers  of  the  Station  for  Experimental  Evolution,  No.  10. 


Munder-Thomsen  Press 
baltimore 


CONTENTS. 


A.  Statement  of  Problem 5 

B.  Material  and  Methods 8 

C.  Results 8 

Series  I.  The  Inheritance  of  Crest 8 

Statement  of  Problem 8 

Material 8 

Results 8 

1.  Recessiveness  of  Plain  Head 9 

2.  The  Detection  of  Homozygous  Crests  and  the  Gametic  Com- 

position of  Heterozygotes 9 

3.  Baldness 11 

Series  H.  The  Inheritance  of  Plumage-Color 13 

Statement  of  Problem 13 

Results 14 

1.  On  Inheritance  of  Green  and  Yellow  Plumage-Color 14 

a.  Green  x  Green 15 

b.  Yellow  X  Yellow 15 

c.  Yellow  X  Green    . 15 

2.  Inheritance  of  Ticking 17 

3.  Inheritance  of  Yellow  x  Lizard  Coloration 19 

4.  Inheritance  of  Cinnamon  (Female)  x  Green  (Male) 19 

Series  III.   Inheritance  of  Characteristics  in  Hybrids  between  the  European 

Goldfinch  and  the  Yellow  Canary 19 

Series  IV.  Hybrids  between  the  Yellow  Canary  and  Other  Species  ...  22 

D.  Summary  and  Conclusion 23 

E.  Table  of  Canary  Matings 24-25 

F.  Literature  Cited 26 

3 


INHERITANCE   IN   CANARIES. 


A.     STATEMENT   OF    PROBLEM. 

The  objection  has  been  raised  that  much  of  the  material  used  in 
studies  of  heredity  has  been  long  under  domestication,  in  consequence 
of  which,  first,  extensive  hybridization  has  occurred,  and,  secondly, 
characteristics  of  an  "abnormal"  sort  have  been  preserved,  and,  as 
a  result,  conclusions  drawn  from  such  material  can  not  properly  be 
applied  to  feral  species  as  they  are  evolving  "in  nature."  It  is 
extremely  doubtful  if  this  objection  has  any  validity,  as  I  have  argued 
elsewhere  (1906).  Nevertheless  it  is  well  to  study  heredity  widely  and 
to  include  in  the  study  some  undomesticated  and  semi-domesticated 
species.  On  this  account,  four  years  ago  I  began  the  breeding  of  certain 
cage-birds  and  especially  the  canary  bird  {Sermus  canarius). 

The  canary  is,  it  is  true,  a  semi-domesticated  animal.  I  say  semi- 
domesticated,  for  in  domestication  there  are  all  degrees.  The  essence 
of  domestication  from  the  standpoint  of  heredity  is  long-continued 
control  by  man  of  mating.  Many  species  of  birds  have  been  bred 
in  zoological  gardens,  and  various  finches — linnets,  siskins,  goldfinches, 
bullfinches,  etc. — are  bred  in  confinement  by  fanciers  and  for  commer- 
cial purposes.  Likewise  grouse  and  quail  and  numerous  species  of 
swimming  birds  have  been  kept  captive  through  many  generations, 
yet  such  acclimated  animals  are  ordinarily  not  regarded  as  domesti- 
cated, because  the  breeding  has  not  been  long  enough  continued  nor 
rigidly  enough  controlled  to  produce  a  number  of  varieties.  Between 
such  acclimated  wild  species  and  thoroughly  domesticated  species,  such 
as  poultry,  dogs,  sheep,  and  horses,  which  are  hardly  to  be  found  in 
the  wild  state,  the  canary  occupies  an  intermediate  position. 

The  history  of  the  canary  in  captivity  is  given  as  follows:*  It 
occurs  as  a  wild  species  in  the  Canary  and  the  Madeira  Islands.  The 
wild  species  agrees  in  coloration  so  completely  with  the  captive  '  'green' ' 
canary  with  greenish-yellow  breast  and  without  white  on  remiges  and 
rectrices  that  only  a  person  with  precise  and  thorough  knowledge  of 
both  can  find  any  certain  distinguishing  characteristic.  After  the 
Spaniards  took  possession  of  the  Canary  Islands  in  1478  they  brought 
many  of  the  products  of  the  islands  home,  and  among  others  these 
remarkable  songsters;  but  for  a  long  time  they  were  rare  in  Europe. 

*The  history  has  been  written  by  Bolle.  e 


6  INHERITANCE    IN    CANARIES. 

Konrad  Gesner,  who  mentions  the  bird  in  his  book  "  De  Avium 
Natura,"  of  which  the  first  edition  was  published  in  1555,  had  never 
seen  it.  According  to  Olina  (1622)  only  males  were,  for  many  years, 
imported,  but  in  the  middle  of  the  sixteenth  century  a  Spanish  ship, 
which  presumably  carried  also  some  female  canaries,  was  wrecked  near 
the  Isle  of  Elba,  and  the  birds  escaped,  and  populated  it  and  created 
there  a  peculiar  strain  in  that  they  were  yellower  beneath  the  chin  than 
those  brought  directly  from  the  Canary  Islands.  These  birds  of  Elba 
were  trapped  by  the  Italians,  bred  in  captivity,  and  sold  in  Italy,  in  the 
Tyrol,  and  in  Germany,  in  which  latter  country  they  were  already  being 
bred  in  captivity  in  many  places  in  the  middle  of  the  seventeenth  cen- 
tury. Already  at  the  beginning  of  that  century  Aldrovandi  had  been 
able  to  get  the  bird  as  a  basis  for  a  figure  in  his  Ornithologiae  (1599-1603). 
From  Germany  the  canaries  were,  in  the  middle  of  the  seventeenth  cen- 
tury, exported  to  England  and  other  countries  and  were  already  regarded 
as  improved  over  the  wild  species,*  but  their  color  was  still  green. t 

The  spread  of  canary  culture  was  aided  by  the  fact  that  they 
became  a  society  fad.  Owing  to  their  high  price,  they  were  attain- 
able only  by  the  wealthy  and  so  became  a  mark  of  that  class.  Ladies 
received  visitors  with  a  canary  perched  on  the  index  finger  and  were 
painted  in  that  attitude,  t 

From  the  beginning  of  the  eighteenth  century  there  is  a  constantly 
increasing  output  of  books  devoted  to  cage-birds  in  general  and  canary 
birds  in  particular,  §  so  that  it  is  possible  to  reconstruct  their  history. 

*InWilloughby's  Ornithology  (Ray,  1678),  quoting  an  earlier  English  author,  it  is 
stated  :  "Canary  birds  of  late  years  have  been  brought  abundantly  out  of  Germany 
and  are  therefore  now  called  German  birds ;  and  these  Germafi  birds  in  handsome- 
ness and  song  excel  those  brought  out  of  the  Canaries. 

tin  "The  Gentleman's  Recreation,"  published  in  1677,  we  find  that  at  that  date 
canaries  in  England  were  of  a  green  color.  (See  Blakston,  Swaysland  &:  Wiener, 
1880,  5.) 

Jin  the  New  York  Public  Library  is  a  little  sociological  tract  entitled,  "Canary 
Birds  Naturalized  in  Utopia  :  A  Canto."     London,  ca.  1708.     The  canto  begins  : 

In  our  unhappy  days  of  Yore, 

When  foreign  Birds,  from  German  Shore, 

Came  flocking  to  Utopia's  Coast, 

And  o'er  the  Country  rul'd  the  Roast :  — 

Of  our  good  People  did  two-thirds 

So  much  admire  Canary  Birds 

For  outward  Show,  or  finer  Feathers 

Far  more  regarded  than  all  others. 

We  bought  'em  dear  and  fed  'em  well, 

Till  they  began  for  to  rebel. 

§The  most  famous  of  early  works  is  that  of  J.  C.  Hervieux:  "Nouveau  traite  des 
Serins  de  Canarie,  contenant  la  maniere  de  les  &  elever  les  appareiller  pour  en  avoir 
de  belles  races ;  avec  des  remarques  aussi  curieuses  que  necessaires  sur  les  signes 
et  causes  de  leur  maladies  et  les  secrets  pour  les  guerir."  12  mo.,  Paris,  1st  edition, 
1705  [Seditions  to  1785].    This  work  was  translated  into  English,  German,  and  Italian. 


Serin  blond  dore 

Serin  blond  aux  duvets 

Serin  jaune  commun 

Serin  jaune  aux  duvets  (race  de  Panaches) 

Serin  jaune  a  queue  blanche  (idem) 


STATEMENT   OF    PROBLEM.  7 

First  it  appears  that  although  in  1677  only  green  canaries  were 
known,  as  early  as  1713  three  main  color  varieties  had  become  estab- 
lished, in  which  various  subvarieties  are  recognized.  The  whole  series 
as  given  by  Hervieux  is  as  follows  : 

Serin  gris 

Serin  gris  aux  pattes  blanches 

Serin  gris  a  queue  blanche 

Serin  blond  commun 

Serin  aux  yeux  rouges 

Hervieux' s  "Serin  gris"  is  doubtless  of  the  wild  type  of  coloration; 
our  "green."  The  "jaune"  is  doubtless  the  modern  clear  or  yellow 
type.  Hervieux  states  that  it  is  (in  1713)  among  the  rare  varieties. 
What  the  "blond"  type  is  can  only  be  conjectured,  but  it  is  probably 
our  mealy,  or  light,  yellow.  If  this  inference  is  correct  three  of  the 
modern  colors  of  canaries — green,  jonque  yellow,  and  mealy  yellow — 
made  their  appearance  at  about  the  end  of  the  seventeenth  century. 
Their  appearance  would  seem  to  have  been  quite  sudden — a  result 
indicating  their  origin  by  mutation  rather  than  by  slow  increments  in 
the  desired  direction.* 

It  appears  also  that  mottled  or  variegated  birds  (race  de  Panaches) 
were  known,  and  as  we  shall  see  later  they  are  probably  the  result  of 
the  crossing  of  a  green  bird  and  a  yellow  one.  Such  variegated  birds 
were  much  esteemed  in  the  early  part  of  the  eighteenth  century. 
During  the  first  half  of  that  century  the  number  of  color  varieties 
was  greatly  increased,  since  Hervieux,  in  his  edition  of  1766,  recorded 
29  color  varieties  including  gris  (green),  blond  (mealy  yellow),  jaune 
(jonque  yellow),  agate,  Isabella  (buff  or  cinnamon),  blanc  (white), 
panache  (mottled),  and  plein — "qui  est  a  present  le  plus  rare." 

The  histories  do  not  state  when  the  crested  form  first  appeared. 
Crested  birds,  like  yellow  ones,  are  now  bred  in  captivity  in  their  native 
islands.  The  frizzled  characteristic  found  in  the  Parisian  Trumpeter 
or  Serin  frise  or  Dutch  frizzle  is  probably  relatively  recent,  as  it  is  rela- 
tively uncommon.  Likewise  of  the  other  varieties  (lizards,  albinos,  etc.), 
the  origin  is  quite  obscure.  The  introducer  of  a  new  variety  usually 
conceals  its  origin;  indeed,  he  has  little  to  say,  as  he  does  not  produce 
or  induce  the  new  characteristic,  but  merely  preserves  that  with  which 
fortune  has  favored  him. 

The  history  of  acclimated  canary  birds  thus  reveals  their  com- 
paratively recent  domestication  and  justifies  the  contention  that  their 
characteristics  may  well  be  expected  to  be  inherited  much  as  they 
would  be  if  found  in  wild  birds. 

The  specific  characteristics  upon  which  I  propose  to  report  in  the 
present  paper  are  two;  viz.,  plumage-color  and  crest.     The  matters  of 


*Russ  (1906,  6)  concludes   that   the    change    from    green    to   yellow   certainly 
occurred  quickly. 


8  INHERITANCE    IN    CANARIES. 

form  and  position,  of  frizzling,  and  certain  others  are  not  yet  ready  for 
a  report.  The  types  of  color  that  I  have  studied  most  thoroughly  are 
the  original  green  and  the  yellow.  These  have  been  bred  pure  i7iter  se, 
and  with  other  species.  The  cinnamon  and  lizard-color  types  have  been 
merely  touched.  As  for  the  crest,  this  consists  of  a  group  of  feathers 
on  top  of  the  head  arranged  so  as  to  radiate  from  a  center  between 
the  eyes  and  forming  a  small  cap  covering  over  the  eyes  and  beak. 
The  questions  are:  How  is  the  crest  inherited,  and  how  is  the  plumage- 
color  inherited  ? 

B.     MATERIAL   AND    METHODS. 

My  original  stock  (1904)  consisted  of  4  yellow  hen  canaries  (one 
crested)  of  the  short  or  German  (Harz  Mountain)  type  and  2  green 
birds  of  the  same  type.  Also  3  yellow  cocks  (one  crested)  and  2 
greens  (one  crested).  To  these  each  year  sundry  other  canaries  were 
added.  These  were  purchased  from  a  dealer  in  New  York  and  had 
been  imported  from  Germany  or  England. 

The  canaries  were  bred  in  a  room  at  the  Station  for  Experimental 
Evolution,  each  pair  being  kept  in  a  separate  cage.  The  usual  methods 
of  feeding  were  adopted.  All  birds  bear  numbered  leg-bands  by  which 
their  identity  is  established. 

C.     RESULTS. 

Series  I.— THE   INHERITANCE   OF   CREST. 
STATEMENT   OF  PROBLEM. 

It  is  my  experience  with  poultry  (1906,  69)  and  pigeons  that  crest 
is  alternative  in  heredity  and  dominates  absence  of  crest.  I  wished  to 
test  heredity  of  crest  in  canaries  also  to  see  whether  the  conclusion  that 
I  have  elsewhere  (1906,  87)  reached  holds,  viz.,  "dominance  and  reces- 
siveness  depend  upon  a  relation  of  the  characteristics  per  se  and  not 
upon  any  relation  of  the  races  into  which  they  have  been  introduced." 

MATERIAL. 

Crested  and  plain  Harz  canaries  were  chiefly  used,  but  also  a  few  of 
the  "Norwich"  type,  which  had  a  flatter  and  darker  crest  (plate  l.fig.  2). 

RESULTS. 

It  quickly  appeared  that  crest  is  alternative  in  inheritance,  for  when 
crested  and  non-crested  birds  were  paired,  the  offspring  were  either 
well  crested  or  plain-headed  and  there  were  no  intergrades.  This  leads 
to  the  hypothesis  that  crest  is  dominant  as  in  poultry  and  pigeons.  To 
test  this  hypothesis  I  made  a  number  of  matings,  of  which  the  detail  is 
given  in  Section  E. 


INHERITANCE    OF    CREST. 


1.  Recessiveness  of  Plain  Head. 

The  following  table  contains,  extracted  from  the  general  table, 
those  experiments  that  give  an  answer  to  the  question  whether  non- 
crested  heads  are  recessive  to  plain  heads.     All  parents  are  non-crested. 

Table  1 .  — Progeny  of  Non- Crested  Parents. 

[The  superior  and  inferior  letters  C  (crested)  and  c  (non-crested)  indicate  the  condition  of  the 
grandparents,  o  signifies  original  stock,  of  whose  ancestry,  consequently,  nothing  is  known  directly. 
The  numbers  in  columns  "Father"   and  "Mother"  are  tliose  of  the  leg-bands.] 


Crest. 

Crest. 

Experi- 
ment No. 

Mother. 

Father. 

F.xperi- 
ment  No. 

Mother. 

Father, 

Absent. 

Present. 

Absent. 

Present. 

401 

7o 

.^O 

1 

0 

618 

K 

151 

8 

0 

404 

/7o 

/6o 

2 

0 

619 

64^ 

ss'i 

11 

0 

405 

/9o 

20o 

2 

0 

623 

^^?. 

K 

8 

0 

501\ 
514/ 

4o 

K 

11 

0 

624 

7Sl 

K 

3 

0 

507 

ISo 

46o 

1 

0 

711 

152\ 

ml 

2 

0 

512 

lOlo 

So 

3 

0 

716 

210\ 

IS5^^ 

3 

» 

605 

S9l 

So 

9 

0 

718 

132\ 

20l\ 

2 

0 

609 

84^ 

^^C 

8 

0 

719 

100\ 

435^^ 

2 

0 

612 

K 

K 

1 

0 

721 

339o 

209\ 

8 

0 

613 

7o 

39^ 

1 

0 

723 

200\ 

1421 

5 

0 

614 
615 

53% 

2So 

8 
2 

0 
0 

725 

412^ 

405\ 

1 

0 

Total  .    . 

.    .   .   . 

102 

0 

Thus  of  102  offspring  of  two  non-crested  parents  all  were  non- 
crested.  In  table  1  are  included  several  cases — Experiments  605,  609, 
619,  623,  and  725 — where  from  one-fourth  to  three-fourths  of  the  grand- 
parentage  is  crested.  In  these  experiments  Galton's  law  calls  for  an 
average  of  ^/  /cast  22.5  per  cent  (and  at  most  45  per  cent)  crested 
offspring.  The  37  offspring  are  all  7ion-crestcd.  Galton's  law  simply 
does  not  apply  to  cases  of  alternative  inheritance. 

2.  The  Detection  of  Homozygous  Crests  and  the  Gametic  Compo- 
sition OF  Heterozygotes. 

If  crest  is  an  alternative  characteristic  we  should  expect  to  find 
some  (one  in  three)  homozygous  dominants  which  always  throw  only 
crested  birds,  whether  mated  with  a  crested  or  a  non-crested  bird.  The 
following  experiments  were  arranged  to  test  the  purity  of  crested  birds. 


10 


INHERITANCE    IN    CANARIES. 


Table  2.  —  To  Test  the  Purity  of  Crested  Parents. 


[I<e!?-1)riii<l  numbers  in  italics  indicate  non-crested  birds.    Arbitrary  signs  employed  in  this  table 
have  the  same  meaning  as  in  table  1.] 


I 

BOTH  PARENTS  CRESTED. 

II. 

ONI,Y  ONE  PARENT  CRESTED. 

Parentage. 

Crest. 

Parentage. 

Crest. 

Isxperi- 
nient  No. 

Experi- 
ment No. 

Mother. 

Father. 

Absent. 

Present. 

Mother. 

Father. 

Absent. 

Present. 

403 

21o 

13o 

1 

1 

402 

Wo 

9o 

0 

1 

504  1 

510/ 

506 

5081 

515/ 

601 

6o 

9o 

1 

10 

502 

44?- 

J60 

2 

5 

12o 
llo 

12o 

13o 
13o 

73^ 
9o 

6,^ 

0 
1 

0 

2 
3 

7 

5031 
511  i 
5051 
515/ 

3^ 

7o 

34C 

4 
3 

5 
5 

608 
610 

67^ 
54C 

0 
0 

5 

2 

514 
604 

]9o 

50o 
62^ 

2 
7 

2 
1 

611 

617 

620 

621 

701 
702 
703 

710 

710a 

713 

97C 

82c 
69^ 

81^ 
207C 
— 0 
67^ 
97C 

I64C 
157^ 

34C 

61^ 

269o 
166^ 

34C 

118^ 

240C 

3 

0 
3 

1 

3 

1 
3 

0 

0 

1 

0 

4 

2 

5 

1 
1 
7 

3 

2 
6 

606 
613 
625 

704 

705 

706 

707 

708 

709 

712 
717 

33o 

J9o 

I27C 

126^ 

I3S^ 

111^ 

178^ 
— 0 

68c 

5O0 

87C 

79C 

125^ 

214^ 

206^ 

73^ 

— 0 

9 

7 

3 

0 

3 

0 

1 

2 

4 

4 
1 

1 

1 

1 

7 

5 

4 

1 

4 

0 

4 
1 

18 

61 

720 

193^ 

1691 

0 

1 

52 

49 

Inspection  of  table  2  shows  that  at  least  2  birds  are  homozygous  in 
crest,  viz.,  No  12  (female),  which  has  produced  9  young,  all  crested, 
and  No.  79  (male),  which  has  produced  11  young,  all  crested.  Also 
No.  126  (female)  (Experiment  706)  is  possibly  homozygous.  No.  9 
(male)  has  produced  an  unexpectedly  large  proportion  of  crested,  viz., 
16  :  1,  where  12.75  :  4.25  was  to  be  expected  ;  mated  with  No.  6  (female). 
No.  79  (male)  was  produced  which  is,  as  just  stated,  homozygous. 

When  two  heterozygous  crested  (DR)  parents  arc  mated  25  per 
cent  of  the  offspring  should  be  non-crested.     Eliminating  the  offspring 


INHERITANCE    OF   CREST.  11 

of  Experiments  506,  601,  and  617  (with  one  parent  homozygous)  we 
have  totals  of  18  non-crested  to  48  crested  offspring,  while  theory- 
calls  in  this  case  for  16.50  and  49.50  respectively.  The  actual  result 
accords  closely  with  expectation. 

When  a  heterozygous  crested  (DR)  parent  is  mated  with  a  non- 
crested  (R)  one,  50  per  cent  of  the  offspring  should  be  non-crested. 
Subtracting  from  the  totals  of  the  right  half  of  table  2  the  offspring 
of  homozygous  79  (male.  Experiment  704),  we  get  52  non-crested  to 
42  crested,  instead  of  the  expected  47  and  47.  The  considerable 
deviation  from  expectation  is  doubtless  due  to  insufficient  numbers 
and  the  extraordinary  run  of  excess  of  non-crested  offspring  obtained 
in  Experiments  604,  606,  and  613. 

3.  Baldness. 

It  is  frequently  stated  (Kidd,  teste  Darwin,  1876,  chap,  viii ; 
Russ,  1906,  127-128)  that  two  crested  birds  should  not  be  mated 
together  because  they  rarely  produce  fine  crests,  but,  on  the  con- 
trary, bald  heads.  One  is  advised  to  mate  the  crested  bird  with  a 
non-crested  one  whose  parents  were  crested.  Blakston  (1880,  129). 
while  approving  this  rule  of  breeding,  scouts  the  idea  that  the  produce 
of  two  crests  tend  to  be  bald.  Such  produce  tend  merely  to  have  high 
tufts  instead  of  the  desired  flat  crests.  The  experiments  listed  in 
table  2  are  of  a  sort  to  throw  light  on  this  question.  Are  the  61  crests 
derived  from  the  experiments  recorded  on  the  left  hand  of  the  table 
different  from  those  derived  from  Experiments  Nos.  505,  515,  604, 
704  to  708,  and  712,  recorded  on  the  right  hand  of  the  table  ?  An 
examination  of  the  distribution  of  bald  offspring  does  not  show  that 
they  are  confined  to  or  exceptionally  prevalent  in  the  cases  of  matings 
of  two  crested  individuals.  It  is  necessary  to  seek  another  cause  for 
baldness. 

An  inspection  of  the  records  shows :  first,  that  of  my  original 
crested  birds  some  had  a  perfect  (or  fully  feathered)  crest  and  in 
others  the  crest  was  imperfect  {i.  e.,  more  or  less  bald  on  the  occiput). 
Birds  with  perfect  crest  were  Nos.  9  (male),  11  (female),  and  34  (male), 
while  birds  with  imperfect  crest  were  Nos.  3  (female),  6  (female), 
12  (female),  13  (male),  and  21  (female).  Now,  with  one  exception  (see 
Experiment  714),  all  of  the  bald  offspring  obtained  in  these  experi- 
ments had  one  of  the  imperfectly  crested  birds  in  its  ancestry.  This 
fact  suggests  the  hypothesis  that  imperfect  crest  is  a  unit-character  in 
heredity.  To  test  this  hypothesis  I  have  made  various  matings  between 
individuals  with  and  those  without  an  imperfect  crest.  The  results  of 
such  matings  are  given  in  table  3. 


12 


INHERITANCE    IN    CANARIES. 
Table  3. — Matings  to  Test  Inheritattce  of  Imperfect  Crest. 


A.     BOTH  PARENTS  HAVE  IMPERFECT  CREST. 
(RRX  RR). 

Experi- 
ment 
No. 

Mother. 

Father. 

Offspring's  Crest. 

Non- 
Crested. 

No. 

Perfect  (P) 
or  Imper- 
fect (I) 
Crest. 

No. 

Perfect  (P) 
or  Imper- 
fect (I) 
Crest. 

Perfect. 

Imperfect. 

506 
617 
620 

12 
84 
69 

I 
I 
I 

13 

79 
83 

I 
I 
I 

0 
0 
0 

2 
1 

1 

'    "    3  "    " 

Total 

0 

4 

3 

B.    NEITHER  PARENT  HAS  IMPERFECT  CREST, 
(a)  DD  X  DD  or  DD  x  DR. 

505 
610 
613 
625 
713 

7 

54 

55 

33 

178 

•     •  p    •     • 
■     ■  p    ■     ■ 

34 
61 

50 

87 

107 

P 
P 
P 
P 

4 
2 
1 
1 
4 

0 
0 
0 
0 
0 

4 

'    '    7  " 
o 

4 

Total  .    . 

12 

0 

18 

(^)    DRXDR. 

705 
707 
70S 
714 

*  127 

*  74 
13S 
157 

P 
•     •  p    •     • 

*  125 
*206 

*  73, 
240 

P 
P 
P 

5 
1 
1 
4 

• 

2 

0 
1 
1 

3 
1 
2 

1 

Total  .    . 

11 

4 

7 

C.     ONE  PARENT  HAS  IMPERFECT  CREST, 
(a)    DR  X  RR. 

508 
510 
515 
603 
604 
608 
703 
704 
706 
711 

11 

6 

11 

11 

19 

67 

67 

19 

126 

164 

P 

I 

P 
P 

'     "    I     "     " 

I 

'     "   I     "     ' 
I 

13 

9 

13 

80 

62 

9 

166 

79 

214 

118 

I 
P 

I 

I 

I 

P 
P 

I 

•     ■  p    ■     ■ 

1 
0 
0 
2 
1 
2 
3 
5 
1 
1 

1 

4 
1 

1 
1 
2 
1 

2 

3 

1 

'    '    3  "    ' 

Total  .    . 

16 

17 

10 

(/3)    DD  X  RR. 

504 
513 
622 

720 

6 

7 

81 

193 

P 
P 
I 
I 

9 

34 

58 

169 

I 
I 
P 

2 
1 
4 
2 

0 
0 
0 
0 

1 

Total  .    . 

9 

0 

1 

*  These  birds  are  known  to  have  imperfect-crested  ancestors  ;  the  others  probably  have. 
Italic  numbers  are  those  of  non-crested  birds. 

INHERITANCE    OF    PLUMAGE-COLOR.  13 

Table  3  shows  that  when  two  imperfect  crests  are  mated  the  off- 
spring have  imperfect  crests  (table  3,  A).  This  indicates  that  absence 
of  occipital  feathering  is  recessive  (R)  to  its  presence.  When  perfectly 
feathered  individuals  that  are  probably  hybrids  between  dominants  (D) 
and  recessives  (R),  i.e.  DR's,  are  mated  with  imperfectly  feathered  con- 
sorts (RR)  an  equal  number  (16  :  17)  of  perfectly  crested  and  imperfectly 
crested  are  thrown  (table  3,  C,  a).  When  two  heterozygous  individuals 
are  mated  there  result  about  75  per  cent  perfectly  crested  offspring  to 
25  per  cent  imperfectly  crested — actually  11:4  (table  3,  B,  /S).  When 
a  pure  dominant  is  mated  to  a  pure  recessive  all  offspring  are  perfectly 
crested  (table  3,  C,  ^).  When  both  parents  have  a  perfect  crest  (being, 
therefore,  DD  or  DR)  all  offspring  have  the  perfect  crest  (table  3,  B,  «). 
Altogether  it  appears  that,  m  crested  birds,  absence  of  feathers  on  the 
occiput  is  recessive  to  their  presence.  Consequently  we  have  in  the 
crest  two  pairs  of  allelomorphs:  crest  6"and  absence  of  crests:  occipital 
feathering  0  and  absence  of  occipital  feathering  (baldness)  o.  Then  CO 
is  the  zygotic  formula  for  perfectly  crested,  Co  for  imperfectly  crested, 
cO  for  the  ordinary  plain-head,  while  a  fourth  possible  combination,  co, 
should  occur  in  one-sixteenth  of  the  offspring  in  F2  and  should  be 
partly  bald  plain-heads.  This  combination  I  seem  not  yet  to  have 
acquired  nor  have  I  seen  it  mentioned  ;  yet  it  is  to  be  expected. 
Nevertheless  it  is  possible  that  baldness  is  coupled  with  crest. 

The  Mendelian  nature  of  the  inheritance  of  baldness  sufficiently 
explains  the  view  of  some  canary  breeders — cited  at  the  beginning  of 
this  section.  For  two  crested  parents  may  throw  from  100  per  cent 
to  25  per  cent  bald  offspring  according  as  they  are  RR  or  DR  in 
respect  to  baldness.  But,  since  all  crestless  parents  are  DD  or  DR 
in  respect  to  baldness,  a  crested  bird  mated  with  a  crestless  can  give 
at  most  only  50  per  cent  bald-headed  and  may  give  none  at  all. 

Series  IL— THE    INHERITANCE    OF    PLUMAGE-COLOR. 
STATEMENT   OF   PROBLEM. 

We  have  seen  that  the  original  canary  was  "green"  and  that  out 
of  such  a  pigmented  canary  there  was  evolved,  over  two  centuries  ago, 
a  yellow  race.  Although  the  historical  evidence  is  incomplete  it  favors 
the  view  that  the  yellow  form  arose  suddenly,  as  a  sport.  Similarly, 
within  recent  years,  in  the  Australian  grass  parakeet,  or  budgerigar,  a 
yellow  variety  has  arisen  under  domestication.  The  interpretation  of 
these  color  changes  must  rest  on  the  facts  of  chemical  physiology. 
According  to  Krukenberg  (1882,  21)  there  is  no  green  pigment  in  the 
wild  canary,  but  the  green  is  due  to  a  yellow  and  a  dark  pigment.  The 
yellow  pigment  is  of  a  fatty  nature,  is  easily  extracted   by  boiling 


14  INHERITANCE    IN    CANARIES. 

alcohol  or  ether,  and  is  classified  as  a  lipochrome  (zoofulvin).  The 
"green"  barbs  from  which  the  lipochrome  has  been  extracted  are 
dark-brown  or  black.  This  dark  pigment  is  much  more  resistant  to 
reagents  and  is  a  melanin  having  its  locus  in  minute  granules.  It  was 
the  purpose  of  the  hybridization  experiments  to  learn  how  these  two 
kinds  of  pigments  are  inherited. 

In  addition  to  green  and  yellow  canaries  there  is  a  color  variety 
known  as  "cinnamon."  A  microscopic  examination  of  the  barb  shows 
that  the  quantity  of  pigment  in  the  quill  feathers  of  the  cinnamon  is 
much  less  than  in  the  "green" — and  is  probably  of  a  different  quality, 
being  allied  to  the  buff  of  pigeons  and  poultry.  A  fourth  color  type 
is  the  "lizard."  This  is  a  special  pattern  rather  than  color,  to  which 
further  reference  is  made  below. 

A  word  about  nomenclature  is  necessary,  and  in  this  I  follow  mainly 
Blakston.  There  are  two  grades  of  concentration  of  all  colors — the 
dense,  bright  shade  or  "jonque"  color,  and  the  dilute,  dull  tint  or 
"mealy"  color.  A  "self"  yellow,  green,  or  cinnamon  is  of  approxi- 
mately one  color  all  over — though  with  varying  shades  and  tints.  A 
yellow  mixed  with  black  over  a  varying  extent  of  the  body  is  called 
mottled  or  pied.  "Heavily  variegated"  birds  have  breast,  back, 
wings,  and  tail  dark;  "lightly  variegated"  birds  have  only  back, 
wings,  and  tail  dark;  "marked"  birds  have  wings  and  tail  dark  and 
no  melanic  pigment  on  the  body  excepting  eye-stripes;  "ticked" 
birds  are  all  yellow,  except  for  small  patches  of  dark  color. 

RESULTS. 

1.  On  Inheritance  of  Green  and  Yellow  Plumage-Color. 

Breeders  of  canaries  for  color  have  formulated  certain  directions 

to  be  followed  in  the  work.     Those  of  Russ  (1906,  125-127),  one  of  the 

best-known  authors  on  cage-birds,  are,  on  account  of  their  definiteness, 

worthy  of  translation  here. 

Experienced  breeders  have  demonstrated  that  in  color-breeding  the  choice  of  the 
male  is  determinative,  while  the  female  must  be  self-colored.  Thus,  e.g.,  to  produce 
crested  offspring  with  a  high  yellow  color  one  selects  a  male  of  this  color,  but  a  green, 
non-crested  female.  The  more  purely  thoroughbred  the  birds  are,  i.  e.,  the  greater  the 
number  of  generations  they  have  been  held  pure,  the  purer  their  progeny.  Let  a 
yellow  pair,  no  matter  of  what  origin,  from  green  or  gray,  produce  young  among  which 
is  a  yellow  male;  let  this  breed  next  year,  in  a  separate  cage,  with  a  yellow  female; 
young  are  produced  among  which  is  a  yellow  male.  If  such  a  male  be  mated  with  a 
female  likewise  bred  true  for  two  generations,  their  progeny  will  never  produce  any- 
thing but  yellow  offspring.  If  one  collects  three  or  four  pairs  of  this  sort  and  lets 
them  fly  free  in  the  room  there  need  be  no  fear  that  any  discolored  birds  will  be 
produced.  .  .  .  The  same  liolds  true  for  all  self-colored  varieties ;  for  the  bright- 
yellows,  straw-yellows,  whites,  greens,  grays,  and  isabella-colored,  but  not  for  mottled 
birds.    The  latter  are  far  more  subject  to  accidental  variation,  and  one  may  be  happy 


INHERITANCE    OF    PLUMAGE-COLOR.  15 

to  obtain  from  four  or  five  broods  one  "Ausstich,"  /.  e.,  a  beautifully  "marked" 
bird.  Here  also  is  it  very  important  that  two  pure-bred  birds  be  mated,  e.  g.,  a 
beautiful  crested  green  cock  with  a  straw-yellow  female;  most  of  the  progeny  will 
then  resemble  the  parents,  and  be  likewise  uniformly  green  or  yellow:  but  if  a  mix- 
ture of  color  does  occur  it  will  usually  be  an  "Ausstich."  Such  "Ausstichvogel" 
are  used  ....  for  the  production  of  rare  markings.  Green  and  Isabella  colors  do 
not  mix,  /.  e.,  these  two  colors  do  not  occur  in  the  same  bird  ;  paired  they  produce 
only  young  which  show  each  color  alone.  Finally,  the  following  results  of  crossing 
seem  to  have  been  demonstrated.  Black  and  green  capped  birds  are  bred  from  a 
male  of  this  sort  mated  with  a  pure  yellow  female;  green  or  black  "swallows"  from 
a  gray  or  blackish-green  crested  male  with  a  non-crested  bright-yellow  female ; 
isabella  "swallows"  from  a  crested  Isabella  male  with  a  golden  yellow  non-crested 
female,  and  gray-,  green-,  and  black-crested  from  a  male  of  the  required  sort  with  a 
bright  or  a  straw-yellow  female. 

My  own  experiments  in  part  confirm  these  statements,  in  part 
disagree  with  them.     These  results  are  as  follows  : 

a.  Green  X  Green. — When  self-greens  are  mated  together  all  of  the 
offspring  are  green.  I  made  five  matings  of  this  sort  (Nos.  402,  404, 
502,  613,  and  624)  and  obtained  21  green  offspring  and  no  other  color. 

b.  YellowX.  Yelloiv. — When  self-yellows  are  mated  together  all  of  the 
offspring  are  yellow.  I  made  11  matings  of  this  sort  and  obtained  34 
offspring ;  all  were  pure  yellow  except  in  6  cases,  where  there  were 
tickings  of  a  darker  color.  These  were  obtained  in  Experiments  Nos. 
503  (2),  513  (1),  and  606  (3).  As  to  Experiment  606  it  should  be  said 
that  No.  68  (male)  used  in  it,  though  yellow,  had  a  green  crest  and  a 
green  patch  lying  off  of  the  crest  on  the  nape  of  the  neck, 

c.  Yellow  Y.  Green. — This  cross,*  no  matter  what  the  ancestry  of 
the  colors,  invariably  gave  mottled  birds  with  varying  proportions  of 
dark  pigment  on   the  yellow  background.     At  one  extreme  are  the 

heavily  variegated  ' '  birds  ;  at  the  other  extreme  is  only  a  ' '  ticking. '  * 
The  usual  result  is  a  "lightly  variegated"  or  unsymmetrically  "marked" 
bird.  The  distribution  of  the  dark  pigment  is  not  wholly  at  haphazard. 
Green  is  usually  found  in  the  lateral  tail  feathers,  on  the  secondaries 
and  wing-coverts  of  one  or  both  wings.  In  addition  to  these  areas 
patches  are  found  on  the  head,  nape,  breast,  at  the  base  of  the  circum- 
anal feathers,  between  the  shoulders,  and  on  the  rump.  The  crossing 
of  yellow  and  green  thus  gave  me,  in  disaccord  with  the  statement  of 
Russ  above,  always  the  mottled  condition  (plate  3,  fig.  5). 

It  now  remained  to  determine  the  nature  of  the  mottled  condi- 
tion— whether  a  fixed  mosaic  or,  like  the  blue  color  of  Andalusian 
fowls,  a  heterozygous  form  or,  like  mottling  in  mice,  due  to  a  mottling 
•factor.  Consequently,  I  mated  the  mottled  birds  of  Fj  with  each  other, 
with  greens,  and  with  yellows.  The  results  of  such  matings  are  given 
in  tables  4  and  5. 

♦Experiments  501,  504,  506,  508,  509,  510,  512,  514,  515,  608,  611,  710. 


16 


INHERITANCE    IN    CANARIES. 
Table  4. — Mottled  X  Mottled. 

[G,  green;  Y,  yellow;  M,  mottled. 1 


Experi- 
ment No. 

Mother. 

Father. 

Offspring. 

No. 

Prevailing 
Color. 

No. 

Prevailing 
Color. 

Yellow. 

Greeu. 

Mottled. 

610 
617 
618 
621 
623 
705 
706 
708 
715 
716 
718 
719 
723 

54 

82 

52 

81 

74 

127 

126 

138 

184 

210 

132 

100 

200 

G 

Y 
Y 
Y 
G 
Y 
Y 
G 
G 
Y 
Y 
Y 
G 

61 

79 
75 

*58 

n 

125 
214 
12> 
103 
186 
201 
135 
142 

G  wings 
G 
Y 
Y 
Y 
G 
Y 
Y 
Y 
Y 
G 
Y 
Y 

0 
0 
3 
1 
0 
5 
1 
0 
0 
0 
0 
0 
3 

1 
1 
0 
0 
0 

1 

0 
2 
0 
0 
0 
0 
1 

1 

0 
5 
4 
8 
2 
3 
2 
1 
3 
2 
2 
1 

13 

6 

34 

*  No.  5Scf  i.s  of  YXY  origin,  but  has  a  green  nape,  perhaps  derived  from  the  grreen  crest  of 
its  mother. 

Table  4  shows  that  mottled  coloration  yields,  upon  inbreeding,  the 
pure  forms,  yellow  and  green. 

Table  ^.—Mottled  X  Self. 

[c^,  male;  ?,  female.] 


MOTTLED  X  GREEN. 

MOTTI^ED  X  YEI.I.OW. 

Mottled  Parent. 

Offspring. 

Mottled  Parent. 

Offspring. 

Experi- 
ment 
No. 

Green 
No. 

Experi- 
ment 
No. 

Yellow 
No. 

No. 

Prevail- 
ing 
Color. 

Mot- 
tled. 

Green. 

No. 

Prevail- 
ing 
Color. 

Mot- 
tled. 

Yellow. 

601 

nc^ 

Y 

12$ 

3 

4 

604 

626" 

Y 

19$ 

6 

4 

603 

SOrT 

G 

11$ 

1 

2 

605 

89$ 

Y 

«rr 

2 

7 

610 

61c^ 

Y  =  G 

54$ 

1 

1 

612 

76$ 

G 

51(i^ 

1 

0 

615 

53? 

G 

28rr 

5 

3 

619 

64$ 

Y 

85d^ 

8 

3 

616 

100? 

G 

37rr 

2 

0 

620 

69$ 

Y  =  G 

83c^> 

3 

1 

624 

78$ 

Y 

37c^ 

1 

2 

702 

269rr 

Y 

Bel.$ 

0 

5 

701 

61d^ 

G 

207$ 

3 

1 

703 

28r^ 

Y 

67$ 

4 

6 

707 

74  9 

G 

206rr 

1 

1 

704 

79c? 

Y 

19$ 

4 

1 

711 

164$ 

G 

118J* 

*1 

0 

709 

111$ 

Y  =  G 

143rr 

4 

0 

714 

240rr 

Y 

157$ 

t4 

2 

720 

193$ 

Y 

169rr 

0 

2 

725 

112$ 

Y 

t05c? 

0 

1 

22 

16 

32 

30 

♦Also  one  yellow.        f  Also  one  yellow  (?)  died  very  young. 


""■"■      'Taleigh 


INHERITANCE    OF   PLUMAGE-COLOR.  17 

The  experiments  recorded  in  table  5  show  that  when  the  mottled 
form  is  mated  with  a  pure  yellow  or  a  pure  green  the  offspring  are 
nearly  equally  of  the  two  parental  colors  and  none  other.*  Moreover, 
the  extracted  yellows  are  "pure,"  since  when  bred  inter  se  they  pro- 
duce only  yellows. 

The  interpretation  of  the  results  of  breeding  plumage-color  is  not 
difficult  and  may  easily  be  brought  to  accord  with  Mendel's  law.  As 
already  stated,  yellow  is  "green"  which  has  lost  its  melanic  pigment. 
The  mottled  canary  further  differs  from  the  green  or  the  yellow  in  a 
spottedness  like  that  of  the  spotted  mouse;  and  as  Cuenot  (1903)  has 
shown  the  latter  to  be  due  to  a  particular  factor  we  may  expect  the 
same  to  be  true  for  the  canary.  Calling  the  black  factor  A'' (nigrum) 
and  the  mottling  factor  M,  we  may  assign  to  the  green  parent  the 
gametic  formula  Nm  and  to  the  yellow  parent  the  formula  7iM,  i.  e.,  it 
contains  the  mottling  factor,  but  lacks  the  melanin  necessary  to  make 
it  show.  The  gametic  composition  of  Fj  is  Wwx"?,  NM  nm  and  the  soma 
shows  black  in  spots  as  a  green  on  a  background  of  yellow.  On  this 
assumption  of  two  pairs  of  allelomorphs,  we  expect  in  every  16  birds 
of  F2  :  9  mottled,  3  green,  and  4  yellow,  of  which  last  class  3  are  uM 
and  1  /iw,  without  trace  of  the  mottling  factor.  The  observed  result 
in  Fg  agrees  fairly  well  with  this  hypothesis.  When  mottled  is  mated 
with  mottled  we  get,  as  table  4  shows,  a  total  of  34  mottled,  6  green, 
and  13  yellow,  expectation  being  30,  10,  and  13  respectively.  The 
result  departs  from  expectation  in  so  far  as  there  is  a  deficiency  of 
greens,  but  a  change  of  4  individuals  from  mottled  to  green  would 
establish  complete  accord  with  theory. 

When  a  mottled  bird  is  mated  with  yellow  or  with  green  expecta- 
tion is  an  equal  number  of  mottled  and  self-colored  offspring.  The 
number  of  offspring  of  each  class  derived  from  the  mottled  X  yellow 
cross  accords  with  expectation.  Of  62,  32  are  mottled  and  30  yellow. 
Of  38  offspring  of  a  mottled  X  green  cross  22,  or  58  per  cent,  were 
mottled  and  16  green.  Here,  again,  is  a  deficiency  of  greens,  but  not 
a  very  improbable  one.  Altogether,  the  results  favor  the  hypothesis 
that  there  are  in  canary  plumage  two  distinct  and  distinguishing 
factors — a  black  factor  and  a  mottling  factor. 

2.  Inheritance  of  Ticking. 

It  has  been  stated  above  that  in  mating  yellow  birds  yellow  off- 
spring ticked  with  black  were  occasionally  obtained.  Similarly  marked 
birds  were  obtained  at  other  times.     It  seemed  desirable  to  ascertain 

*One  clear  exception  and  one  doubtful  one  are  found   in  the  green  X  mottled 
rnatings  of  Experiments  711  and  714  where  a  yellow  appeared  among  the  offspring. 


18 


INHERITANCE    IN    CANARIES. 


whether  this  ticking  is  an  accident  that  may  be  eliminated  by  further 
dilution  with  yellow  blood  or  darkening  with  "green"  blood  or  whether 
it  is  a  unit-character  like  mottling  which  persists,  defying  all  attempts 
at  dilution.  The  history  of  some  matings  to  test  this  point  is  given  in 
table  6,  in  detail. 

Table  6. 


Exp. 
No. 

Mother. 

Father. 

Offspring. 

No. 

From 
Exp. 

No. 

Description. 

No. 

From 
Exp. 
No. 

Description. 

Nos.  and  Description. 

621 

81 

510 

Mottled 

58 

503 

Y  +  N  on 
nape 

127,  129,  189,  mottled. 
188,  Y  +  N  patch  at  base  of  beak. 
190,  Y  +  N  spots  on  head  and 
nape. 

605 

89 

513 

Y  +  N 
patch  on 
L  nape 

8 

o 

Y 

168,  169,  170,  171,  all  Y. 

135,  Y  +  N  spot  on  R  nape  and 
between  shoulders. 

136,  Y  +  N  spot  on  L  wing. 

721 

339 

o 

Y 

209 

618 

Y  +  N 
spot  on 
R  nape 

271,273,  317,  all  Y. 

270,  Y  +  spot  under  L  eye. 

272,  Y  +  neck  stripe,  L  side. 

314,  Y  +  spot  at  L  ear. 

315,  Y  +  spot  over  each  eye,  on 
R  nape  and  R  secondaries. 

316,  Y  +  spot  at  L  eye  and  on  L 
wing. 

Table  6  may  be  summed  up  in  the  statement  that  ticked  yellow 
behaves  like  variegation  ;  for  ticked  yellow  X  clear  yellow  gives  50  per 
cent  ticked  and  50  per  cent  pure  yellow.  Ticking  differs  from  variega- 
tion only  in  the  amount  of  dark  pigments  involved.  We  have  already 
seen  that  there  is  probably  a  determiner  for  mottling.  We  now  see 
that  the  mottling  determiner  occurs  in  various  degrees  which  maybe 
designated  M',  M",  etc. 

The  question  arises  what  determines  the  degree  of  mottling  in  any 
case  ?  If  all  mottling  results  from  a  cross  of  yellow  and  green  why  are 
the  proportions  of  yellow  and  green  so  diverse  ?  Of  the  fact  of  this 
diversity  there  is  no  doubt.  For  example,  in  Experiment  501  a  pure 
yellow  (No.  4  female)  was  crossed  with  a  pure  green  (No.  37  male), 
and  of  two  offspring  one  was  green,  except  for  yellow  bands  across  the 
back  and  a  yellow  belly  and  breast.  The  other  was  all  yelloiv  except 
for  dark  eye-spots,  side  of  breast,  and  base  of  perianal  fluff.  Conse- 
quently one  may  speak  of  the  mottling  factor  as  wide  in  one  case  and 
restricted  in  the  other.  Individual  germ  cells  vary  in  the  extent  of  the 
spots  they  determine. 


INHERITANCE    OF    PLUMAGE-COLOR.  19 

3.  Inheritance  of  Yellow  X  Lizard  Coloration. 

The  "  Lizard  "  canary  is  closely  related  to  the  "Green  ;  "  it  differs 
in  that  the  margin  of  each  of  the  dorsal  body  feathers  is  much  lighter 
than  the  rest  of  the  web — the  feathers  are  "laced."  This  condition 
is  dimly  seen  in  the  green  canary,  but  is  much  exaggerated  in  the 
"Lizard."  The  light  tip  may  be  either  yellow  (jonque)  as  in  the 
"Gold  Lizard"  or  white  (mealy)  as  in  the  "Silver  Lizard."  A  second 
characteristic  of  the  Lizard  is  a  light  area  or  "cap"  on  top  of  the 
head  over  the  eyes,  which,  similarly,  is  either  reddish  or  white.  The 
characters  of  the  Gold  and  of  the  Silver  Lizard  are  said  to  be  quite 
stable  in  straight  breeding. 

In  my  experiments  (Nos.  507  and  512)  I  crossed  a  Gold  Lizard 
(male)  with  a  crestless  yellow  Norwich  canary.  One  young  was 
obtained.  This  had  the  yellow  cap  of  the  Lizard  parent.  The  bird 
was  mottled,  but  the  dark  feathers  were  laced  (like  those  of  the  Gold 
Lizard)  with  yellow.  When  a  Silver  Lizard  (female)  was  mated  with 
a  yellow  Harz  I  got  three  young,  closely  alike.  All  had  a  cap,  but 
this  was  yellow  instead  of  silver.  All  were  mottled  and  the  dark 
body  feathers  were  generally  laced.  It  appears,  therefore,  from  these 
few  experiments,  that  cap  and  lacing  are  dominant  over  their  absence 
and  "gold"  is  dominant  over  silver,  Z.^".,  jonque  over  mealy. 

4.  Inheritance  of  Cinnamon  (Female)  X  Green  (Male). 

In  canary  hybridization  cinnamon  has  long  been  known  to  behave 
in  a  peculiar  way.  A  cinnamon  male  mated  with  a  female  of  another 
race  may  produce  some  cinnamon  offspring;  but  a  pure  non-cinnamon 
male  mated  to  a  cinnamon  female  produces  no  cinnamons.  Also,  the 
offspring  of  cinnamons  bred  to  greens  are  often  of  a  better  cinnamon 
color  than  their  parents.  My  own  experience  consists  of  two  progeny 
of  a  cinnamon  Belgium  female  X  green  Belgian  male.  Both  are  of 
a  beautiful  green  color  and  show  no  trace  of  the  cinnamon  color.  The 
female  cinnamon  seems  to  be  fully  recessive  to  green. 

Series  IIL— INHERITANCE  OF  CHARACTERISTICS  IN  HYBRIDS  BETWEEN 
THE  EUROPEAN  GOLDFINCH  AND  THE  YELLOW  CANARY. 

The  statement  is  frequently  seen  that  it  makes  a  great  difference 
in  heredity  whether  the  individuals  crossed  belong  to  allied  races  or  to 
distinct  species.  Fockc  (1881, 473)  states  that  in  race-hybrids  characters 
of  the  parents  do  not  blend  as  they  do  in  species-hybrids.  It  is 
important  to  know  if  this  law  holds  universally,  and  the  finches  offer 
a  good  opportunity  to  test  it.  They  are  easily  hybridized  and  the 
results  of  such  experiments  have  often  been  recorded  by  writers  on 
cage-birds  {cf.  Blakston).     The  commonest  of   these   hybrids   is   that 


20  INHERITANCE    IN    CANARIES. 

between  the  male  goldfinch  and  female  canary.  These  hybrids  have 
been  often  described  and  they  have  been  carefully  analyzed  by  Klatt 
(1901),  who  used,  however,  only  museum  material  or  descriptions  and 
knew  little  about  the  parents  of  the  individual  birds  examined.  Like 
all  other  writers  on  the  subject,  Klatt  lays  emphasis  on  the  great  vari- 
ability of  the  first  generation  of  hybrids — a  variability  which  is  in 
striking  contrast  to  the  uniformity  exhibited  by  most  first  hybrids 
between  domesticated  races.  The  hybrids  between  the  goldfinch  and 
the  canary  are  usually  very  dark — brown,  black,  and  "green"  pre- 
dominating— but  they  usually  show  various  yellow  and  white  patches 
which  may  be  very  extensive  and,  in  extreme  cases,  result  in  almost 
entire  albinos.     This  variability  demands  an  explanation. 

The  goldfinch  {JFri7igilla  cardnelis  Linnaeus),  as  shown  in  plate  2, 
fig.  4,  is  marked  on  the  head  by  a  red  patch  on  forehead  and  chin,  a 
black  eye-stripe  and  a  black  cap  extending  back  on  to  the  nape,  where 
it  is  sharply  cut  off  by  a  transverse  white  band.  A  pair  of  white  areas 
run  up  from  the  throat  on  the  sides  of  the  head  to  the  black  cap.  On 
the  body,  the  back  and  sides  are  brown,  this  color  extending  also  all 
over  the  breast  and  upper  wing  coverts.  The  rest  of  the  ventral  body 
is  white.  Yellow  areas  on  the  middle  of  the  exposed  portion  of  all 
quill  feathers  form  in  the  folded  wing  a  yellow  wing  band,  and  the 
quill  feathers  are  tipped  with  white. 

When  such  a  goldfinch  was  crossed  with  a  crested  yellow  Harz 
(plate  1,  fig.  2)  one  of  the  hybrids  was  like  plate  2,  fig.  3.  One  can 
see  at  a  glance  that  the  hybrid  is  not  a  mere  combination  of  the 
characters  of  the  two  parental  forms,  but  is  more  like  a  green  canary 
combined  with  the  goldfinch.  First,  a  rudimentary  crest  is  present. 
The  red  of  the  face  has  become  of  a  copper  color  (red  +  yellow)  and 
the  cap  is  dark  greenish  (black  +  yellow).  The  breast  and  belly  are 
yellowish  as  in  the  "green"  canary.  The  remiges  are  black  with 
lighter  tips — a  modified  goldfinch  character.  The  yellow  wing-bar  is 
present,  but  reduced,  combining  the  character  of  the  goldfinch  with 
that  of  the  green  canary.  The  sides  of  breast  and  wing-coverts  are 
striped,  due  to  a  central  blackening  of  the  feathers — a  character  of  the 
green  canary.  It  thus  appears  that  characters  of  the  green  canary 
predominate,  but  do  not  replace  the  more  striking  characters  of  the 
goldfinch. 

The  fact  that  the  hybrids  between  the  goldfinch  and  yellow  canary 
have  many  of  the  distinctive  features  of  the  "green"  canary  has  been 
frequently  observed.  Darwin,  having  heard  of  the  streaked  feathers 
of  the  hybrid,  concluded  that  "this  streaking  must  have  been  derived 
from  the  original  wild  canary;"  and  this  case  seemed  to  favor  his 
theory  of   reversion.     Klatt  (1901,  508)  goes  further  and   concludes 


CHARACTERISTICS    IN    HYBRIDS.  21 

that  there  is  a  reversion  in  some  characters  to  the  serin  or  Girlitz 
{Serifius  hortulatius  Koch),  probably  close  to  the  ancestor  of  the  wild 
canary.  These  characters  are  four:  "in  der  breiten  Binde  der  Spitzen 
der  Armschwingen,  den  Fliigelbinden  der  Deckfedern,  den  Farbung 
der  Schultern,  den  Saumen  der  Steuerfedern."  But  in  respect  to  the 
first  and  last  of  these  characters  I  find  in  my  birds  no  important  differ- 
ence from  the  green  canary,  except  a  slight  yellowing  of  the  lighter 
areas.  The  color  of  the  shoulders  of  the  hybrid  is,  as  it  were,  the 
sum  of  the  colors  of  the  goldfinch  and  the  canary.  As  to  the  second 
of  the  differential  characters— the  yellow  wing-bow — this  is  highly 
variable  in  my  hybrids  and  in  mottled  canaries.  We  have  no  reason  to 
conclude  that  there  is  reversion  to  the  serin,  and  it  is  undesirable  to 
rest  with  so  vague  a  term  as  reversion  as  an  "explanation"  of  the 
resemblance  of  the  hybrid  to  the  "green  canary." 

But  if  not  reversion,  under  what  rubric  shall  we  place  the  greenness 
of  the  goldfinch  X  canary  hybrid  ?  First  it  is  to  be  recalled  that  the 
yellow  canary  is  a  green  deprived  of  black  pigment.  When  black  pig- 
ment is  added  from  any  source  it  occupies  the  emptied  spaces  and  so 
restores  the  "green"  and  the  black.  Consequently  we  find  streaking 
on  the  sides  of  the  body  in  the  goldfinch-canary  hybrid  and  black  on 
tail  and  wings.  But  the  canary  contains  also  a  mottling  factor  and  so 
the  hybrid  is  "green"  in  certain  areas  only.  The  other  areas  are 
yellow  or  else  yellow  to  which  the  chocolate  color  has  been  added  on 
the  back,  wing  coverts,  and  sides,  and  to  which  red  has  been  added  on 
the  face.  The  belly,  which  is  white  in  the  goldfinch,  remains  yellow 
in  the  hybrid. 

How,  then,  shall  we  conceive  the  gametic  formula  of  the  gold- 
finch and  the  yellow  canary  ?  As  the  goldfinch  contains  black  (A^), 
red,  and  chocolate,  its  formula  may  be  given  as  N^R,  C,  m,  while  that 
of  the  canary  is  fi,  r,  c,  M,  contributing  only  the  mottling  character. 
Then  the  zygote  gives  NRCAIas  dominants,  and  the  adult  hybrid  shows, 
on  top  of  the  yellow,  black,  red  on  the  face  and  chocolate  on  the  back 
and  the  sides. 

The  foregoing  theory  of  the  gametic  constitution  of  the  yellow 
helps,  moreover,  to  explain  the  great  variability  of  the  hybrids — corre- 
sponding to  the  variability  of  the  mottled  offspring  of  yellow  X  green. 
As  we  have  seen  that  in  an  extreme  case  the  hybrid  yellow  X  green  is 
practically  yellow,  so  likewise  the  hybrid  yellow  X  goldfinch  is  occa- 
sionally, though  rarely,  entirely  or  almost  entirely  yellow  except  for 
the  red  on  the  face. 

Finally,  attention  must  be  called  to  the  principle  of  localization  of 
unit-characters.  Red  is  found  almost  invariably  in  the  goldfinch 
hybrid,  but  always  confined  to  the  head  region.     Chocolate  occurs  in 


22  INHERITANCE    IN    CANARIES. 

the  hybrid,  if  at  all,  only  where  it  is  present  in  the  goldfinch.  Black, 
on  the  other  hand,  is  a  color  that  belongs  both  to  the  wild  canary  and 
the  goldfinch.  The  yellow  canary  has  merely  lost  one  factor  necessary 
to  the  production  of  black  in  these  situations  where  black  occurs  in  the 
green  canary.  When  black  is  introduced  by  the  goldfinch  it  is  laid 
down  (1)  in  accordance  with  the  goldfinch  formula  and  (2)  also  in 
accordance  with  the  green  canary  formula,  but  (3)  both  distributions 
are  controlled  by  the  variegation  factor,  so  that  black  (like  the  other 
pigments)  is  laid  down  only  over  more  or  less  circumscribed  areas  of 
the  body. 

Series  IV.-^HYBRIDS    BETWEEN    THE   YELLOW    CANARY 
AND   OTHER   SPECIES. 

Canary  fanciers  have  been  very  active  in  making  "mules"  between 
various  species  of  finches.  Besides  the  goldfinch,  the  linnet  {Fri?igilla 
linota),  the  greenfinch  {Fritigilla  chloris),  and  the  siskin  {Frmgilla  spinus) 
have  been  crossed  with  the  canary. 

The  linnet  is  a  prevailingly  brown  bird,  with  black,  white-edged 
quill  feathers  and  darker  striping  on  wing  coverts  and  sides  of  body. 
The  hybrids  with  the  yellow  canary  are  said  usually  to  be  dark  birds 
resembling  the  linnet.  Mottled  birds  sometimes  occur,  linnet  colora- 
tion showing  in  patches  on  the  otherwise  yellow  background.  (Blak- 
ston,  1880,  272).  Here  the  brown  of  the  linnet  seems  to  dominate  over 
the  canary  green,  but  the  mottling  factor  of  the  yellow  canary  is  active. 

The  greenfinch  is  olive-green  above,  yellowish-green  below,  has 
black  remiges  edged  with  yellow  (forming  a  wing-bar)  and  black  rec- 
trices  edged  with  olive-green,  except  the  four  outer  tail  feathers,  which 
are  edged  with  bright  yellow.  The  hybrids  are  sometimes  dark  like 
the  greenfinch,  but  "highly  variegated"  are  common.  Here,  again,  is 
seen  the  mottling  factor  of  the  canary. 

The  siskin  is  streaked  greenish  above  anteriorly  and  yellowish 
posteriorly;  below  is  light  green  in  the  male  and  white  in  the  female. 
There  is  a  yellow  stripe  over  the  eye.  The  sides  of  body  and  wing 
coverts  are  distinctly  striped.  The  wings  and  tail  are  prevailingly 
black,  but  the  base  of  the  rectrices  and  a  wing-bar  and  wing-bow  are 
yellow.  The  hybrid  with  the  yellow  canary  (plate  1,  fig.  2)  closely 
resembles  the  siskin  — the  dark  beak,  the  yellow  supraorbital  stripe, 
the  striping,  even  to  the  lower  tail-coverts,  are  all  present.  In  my 
hybrids  the  wing-coverts  and  rump  are  green  as  in  the  green  canary. 
Here  again  the  hybrid  shows  the  so-called  reversion  phenomena. 
Mottling  occurs  in  the  hybrids  under  certain  conditions,  but  the  usual 
type  is  all  dark  (plate  3,  fig.  6). 

The  foregoing  descriptions  show  first  that  in  all  hybrids  between 
the  yellow  canary  and  a  finch  there  is  a  tendency  to  "reversion" — a 


SUMMARY   AND    CONCLUSION.  23 

result  that  falls  into  the  same  category  as  the  "reversion"  of  the 
goldfinch  hybrid.  In  all  cases  the  hybrids  with  the  yellow  canary 
are  very  variable  and  frequently  show  more  or  less  of  the  canary  yel- 
low. This  is  due  to  the  mottling  factor  of  the  canary  to  which  refer- 
ence has  been  so  often  made.  That  it  is  the  yellow  canary  which 
contains  the  mottling  factor  and  is  the  source  of  the  variability  of  the 
hybrids  is  shown  by  the  facts  that  (1)  hybrids  with  the  green  canary 
do  not  vary  in  this  fashion,  and  (2)  hybrids  between  any  two  species  of 
finches — of  which  many  are  bred  by  fanciers — are  "cast  in  one  mold." 

D.     SUMMARY   AND   CONCLUSION. 

The  history  of  the  domestic  canary  shows  that  it  has  been  intensely 
bred  for  only  about  250  years  and  may  therefore  be  regarded  as  a  rela- 
tively recently  acclimated  species  when  compared  with  poultry  that  have 
been  bred  for  over  2,000  years.  Nevertheless  distinctive  characters  have 
arisen  which  behave  in  Mendelian  fashion. 

Crest  is  dominant  over  plain  head. 

Baldness  is  a  unit-character  and  is  recessive  to  perfect  crest. 

The  yellow  canary  is  derived  from  the  original  "green"  canary 
by  the  loss  of  black.  It  carries  a  mottling  factor.  Consequently  when 
the  yellow  canary  is  crossed  with  a  pigmented  canary  or  with  a  finch 
the  hybrids  are  mottled. 

The  mottling  is  not  a  fixed  pattern.  The  spots  vary  in  position 
and  relative  size — they  may  cover  nearly  the  whole  body  or  they  may 
form  a  mere  "ticking."  The  degree  of  mottling  is  inheritable.  Ticking 
behaves  as  a  unit-character. 

Mottling  is  a  heterozygous  character  and  throws  mottled,  clear 
yellow  and  self-greens. 

The  principle  of  localization  of  the  units  of  a  complex  plumage 
must  be  recognized.  The  cap  of  the  Lizard  canary,  the  red  face  of 
the  goldfinch,  the  shoulder  striping  of  the  green  canary  are  not  only 
unit-characters  but  they  occur  only  at  their  proper  localities  and  in 
their  proper  forms  in  the  body  plumage.  In  mottled  canaries  the 
presence  of  black  on  the  shoulder  means  striping,  on  the  wing  it  means 
dead  black,  white-laced  remiges,  on  the  mid-breast  it  means  a  uniform 
olive  color.  The  plumage  of  a  yellow  canary  may  be  compared  with  a 
letter  that  has  been  written  with  invisible  ink.  Wherever  the  developer 
acts  (i.  e.,  the  black  pigment  of  the  green  canary  is  added)  that  which 
is  written  appears  with  all  of  its  idiosyncrasies. 

Cold  Spring  Harbor,  N.  Y., 
September  28,  1907. 


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25 


26  INHERITANCE    IN    CANARIES. 

F.     LITERATURE   CITED. 

Aldrovandi,  U. 

1599-1603.     Ornithologi^,  etc.     Bononias. 

Anonymous. 

1708  [?].     Canary-Birds  Naturalized  in  Utopia.     A  canto.     London,  n.  d. 

1735.     The  Bird-Fancier's  Recreation:  Being  Curious  Remarks  On  the  Nature 
of  Song-Birds,  etc.     London. 
Blakston,  W.  a. 

1880  [?].  The  Illustrated  Book  of  Canaries  and  Cage-birds.  London.  Casseli, 
n.  d.  [The  Britisli  and  Foreign  Cage-I3irds  by  W.  Swaysland  and 
A.  F.  Wiener,  respectively.] 

CUBNOT,   L. 

1903.     L'heredite  de  la  pigmentation  chez  les  Souris  noires.    Archives  de  zoologic 
exper.  et  gen.  [4],  tom.  i,  Notes  et  Revue,  p.  xxxiii-xii. 

Darwin,  C. 

1876.     The  variation  of  animals  and  plants  under  domestication.     Second  edition, 
revised,  vols.  I,  II.     Ncw^  York :  D.  Appleton  &  Co. 

Davenport,  C.  B. 

1906.     Inheritance  in  Poultry.     Publications  of  the  Carnegie  Institution  of  Wash- 
ington No.  52  (Papers  Station  for  Experimental  Evolution  No.  7). 
1906  «.     The    Mutation   Theory    in   Animal    Evolution.      Science,  n.  s. ,  xxiv, 
pp.  556-558,  Nov.  2,  1906. 

FOCKE,  W.  O. 

1881.  Die  Pflanzen-Mischlinge.     Ein  Beitrag  zur  Biologie  der  Gewachse.     Ber- 

lin: Borntraeger,  iv-|-567  pp. 

Gesner,  C. 

1617.  Historiae  Animalium.  Liber  II,  qui  est  de  Avium  natura,  etc.  Franco- 
furti.     [Edit  1,  Fol.  Figuri,  1555.] 

Godman,  F.  du  C. 

1870.  Natural  History  of  the  Azores  or  Western  Islands.  London:  van  Voorst, 
v-l-358  pp. 

Hervieux  de  Chanteloup,  J.  C. 

1705.  Nouveau  traite  des  serins  de  canarie.  Paris.  [The  reference  in  the  text 
is  from  the  edition  of  1713 — the  2d  edition.  The  Station  for  Experi- 
mental Evolution  has  the  edition  of  1766 — the  fourth.  An  English 
translation  appeared  in  London,  1718.] 

Klatt,  G.  T. 

1901.  tJber  den  Bastard  von  Stieglitz  und  Kanarienvogel.  Arch,  fiir  Entwicke- 
lungsmeclianik  der  Organismcn.  xii.  414-528.  Taf.  ix.  August  2d 
and  September  10th. 

Krukenberg,  C.  F.  W. 

1882.  Vergleicliend-physiologische  Studien.     Zweite  Reihe,  Zweite  Abtheilung. 

Heidelberg:  Winter.     96  pp. 

Noorduijn-Groningen,  C.  L.  W. 

1905.  Die  Farben-  und  Gestaltskanarien.     Magdeburg:  Creutz.     vni+152  pp. 
Olina,  G.  p. 

1622.     Uccelliera,  etc.     Roma. 

Ray,  J. 

1678.  Tlie  Ornithology  of  Francis  Willoughby,  etc.  London  :  Martyn.  442  pp. 
78  tab. 

Russ,  K. 

1906.  Der  Kanarienvogel:  Seine  Naturgeschichte,  Pflege  und  Zucht.     11  Auf. 

Bearbeitet  von  R.  Hoffschildt.     Magdeburg  :  Creutz.     244  pp. 


PLATE  1 


^^ 


■/ 


/ 


FIG.   1.     CRESTED      GREEN"  CANARY   (NO.   1  1  8  a") 

FIG.   2.     CRESTED  YELLOW  CANARY,   MEALY   TYPE   (NO.   67?) 


PLATE  2 


A  MOtM  •  CO  SALT  <  - 


FIG.   3.     HYBRID  BETWEEN  GOLDFINCH   (FIG.  4)   AND  CRESTED  YELLOW  CANARY.     TYPE  OF  FIG.   2   (NO.   203.,') 
FIG.  4.     EUROPEAN  GOLDFINCH,   FATHER  OF   NO.  203   (FIG.  3) 


PLATE  3 


5. 


A  Hvi  iJi  lo  iv^Lnvour 


FIG.   5.     MOTTLED  OFFSPRING  OF  CANARIES  OF  TYPES  OF  FIGURES   1    AND  2.     HIGHLY  VARIEGATED  TYPE   (NO.  61  A 

FIG.  6.     HYBRID  BETWEEN   SISKIN  AND  CRESTED  YELLOW  CAri«|^^(NO.   264) 


CRESTED  YELLOW  CA^r^^KjNO 


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